T Mizuno

Nagoya University, Nagoya, Aichi, Japan

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Publications (301)1011.19 Total impact

  • [Show abstract] [Hide abstract]
    ABSTRACT: During the last decade, significant research progress in Arabidopsis thaliana has been made in defining the molecular mechanisms behind the plant circadian clock. The circadian clock must have the ability to integrate both external light and ambient temperature signals into its transcriptional circuitry to properly regulate its function. We previously showed that transcription of a set of clock genes including LUX (LUX ARRHYTHMO), GI (GIGANTEA), LNK1 (NIGHT LIGHT-INDUCIBLE AND CLOCK-REGULATED GENE 1), PRR9 (PSEUDO-RESPONSE REGULATOR 9) and PRR7 is commonly regulated through the evening complex (EC) nighttime repressor in response to both moderate changes in temperature (Δ6°C) and differences in steady-state growth-compatible temperature (16°C to 28°C). Here, we further show that a nighttime-light signal also feeds into the circadian clock transcriptional circuitry through the EC nighttime repressor, so that the same set of EC target genes is upregulated in response to a nighttime-light pulse. This light-induced event is dependent on phytochromes, but not cryptochromes. Interestingly, both the warm-night and nighttime-light signals negatively modulate the activity of the EC nighttime repressor in a synergistic manner. In other words, an exponential burst of transcription of the EC target genes is observed only when these signals are simultaneously fed into the repressor. Taken together, we propose that the EC nighttime repressor plays a crucial role in modulating the clock transcriptional circuitry to properly keep track of seasonal changes in photo- and thermal-cycles by conservatively double-checking the external light and ambient temperature signals.
    Plant and Cell Physiology 10/2014; · 4.98 Impact Factor
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    ABSTRACT: An interlocking multi-loop model has been generally accepted to describe the transcriptional circuitry of core clock genes, through which robust circadian rhythms are generated in Arabidopsis thaliana. The circadian clock must have the ability to integrate ambient temperature signals into the clock transcriptional circuitry to properly regulate clock function. Clarification of the underlying mechanism is a longstanding subject in the field. Here, we provide evidence that temperature signals feed into the clock transcriptional circuitry through the evening complex (EC) nighttime repressor consisting of ELF3, ELF4, and LUX (also known as PCL1). Chromatin immunoprecipitation assays showed that PRR7, GI and LUX are direct targets of the nighttime repressor. Consequently, transcription of PRR9/PRR7, GI and LUX is commonly regulated through the nighttime repressor in response to both moderate changes in temperature (Δ6°C) and differences in steady-state growth-compatible temperature (16°C to 28°C). A warmer temperature inhibits EC function more, whereas a cooler temperature stimulates it more. Consequently, the expression of these target genes is upregulated in response to a warm temperature specifically during the dark period, whereas they are reversibly downregulated in response to a cool temperature. Transcription of another EC-target, the PIF4 gene, is modulated through the same thermoregulatory mechanism. The last finding revealed the sophisticated physiological mechanism underlying the clock-controlled output pathway, which leads to the PIF4-mediated temperature-adaptive regulation of hypocotyl elongation.
    Plant and Cell Physiology 02/2014; · 4.98 Impact Factor
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    ABSTRACT: Ambient temperature has two fundamental impacts on the Arabidopsis circadian clock system in the processes referred to as temperature compensation and entrainment, respectively. These temperature-related longstanding problems have not yet been fully clarified. Recently, we provided evidence that temperature signals feed into the clock transcriptional circuitry through the evening complex (EC) nighttime repressor composed of LUX-ELF3-ELF4, and that the transcription of PRR9, PRR7, GI and LUX is commonly regulated through the nighttime repressor in response to both moderate changes in temperature (∆6 °C) and differences in steady-state growth-compatible temperature (16 °C to 28 °C). These temperature-associated characteristics of the core clock genes might be relevant to the fundamental oscillator functions. Here, we further show that the recently identified LNK1 night light-inducible and clock-controlled gene, which actually has a robust peak at daytime, is induced also by warm-night through the EC nighttime repressor in a manner very similar to PRR7, which is also night light-inducible daytime gene. Based on these findings, a hypothetical view is proposed with regard to the temperature entrainment of the central oscillator.
    Plant signaling & behavior 01/2014; 9(3).
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    ABSTRACT: Plant elongation growth on a day-to-day basis is enhanced under specific photoperiod and temperature conditions. Circadian clock is involved in the temperature adaptive photoperiodic control of plant architecture, including hypocotyl elongation in Arabidopsis thaliana. In this regulation, phytochrome interacting transcriptional factors, PIF4 and PIF5, are activated at the end of night under short photoperiod or high temperature conditions, due to the coincidence between internal (circadian rhythm of the transcripts) and external (length of dark period) time cues. It is previously found that biosynthesis or metabolism of phytohormones including auxin, and their signal transduction-related genes are downstream targets of circadian clock and PIF4/PIF5 mediated external coincidence mechanism. Brassinosteroid and gibberellic acid played a positive role in the hypocotyl elongation of seedlings under light and dark cycle conditions. On the other hand, cytokinin and jasmonic acid played an opposite role. In this study, diurnal expresson profile of a gene encoding a sulfotransferase family protein that is involved in the jasmonic acid metabolism, ST2A, was examined. It was found that transcription of ST2A is induced at the end of night under LD/22 °C and SD/28 °C conditions according to the external coincidence mechanism. The results of this study support the idea that the circadian clock orchestrates a variety of hormone-signaling pathways to regulate the photoperiod and temperature-dependent morphogenesis in A. thaliana.
    Bioscience Biotechnology and Biochemistry 12/2013; · 1.27 Impact Factor
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    ABSTRACT: Plant circadian clock generates rhythms with a period close to 24 h, and it controls a wide variety of physiological and developmental events, including the transition to reproductive growth (or flowering). During the last decade, significant research progress in Arabidopsis thaliana has been made in defining the molecular mechanism by which the circadian clock regulates flowering time in response to changes in photoperiod. In Lotus japonicus, we have found that LjFTa, which encodes a ortholog of the Arabidopsis FLOWERING LOCUS T (FT), plays an important role in the promotion of flowering, but it is not clear how the expression of LjFTa is regulated in L. japonicus. Based on current knownledge of photoperiodic control of flowering time in A. thaliana, here we examined whether a microRNA is involved in the activation of LjFTa in L. japonicus. Two putative L. japonicus genes that are responsible for the production of miR172 (designated LjmiR172a and LjmiR172b) were cloned. Overexpression of LjmiR172a/b in A. thaliana resulted in markedly accelerated flowering through enhancement of the expression of FT, concomitantly reducing the expression level of TARGET OF EARLY ACTIVATION TAGGED 1 (TOE1) transcripts, the protein product of which functions as a transcriptional repressor of FT. These results suggest that LjmiR172 genes play a positive role in the LjFTa-mediated promotion of flowering in L. japonicus.
    Bioscience Biotechnology and Biochemistry 06/2013; · 1.27 Impact Factor
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    ABSTRACT: During the last decade, significant research progress in the study of Arabidopsis thaliana has been made in defining the molecular mechanism by which the plant circadian clock regulates flowering time in response to changes in photoperiod. It is generally accepted that the clock-controlled CONSTANS (CO)-FLOWERING LOCUS T (FT)-mediated external coincidence mechanism underlying the photoperiodic control of flowering time is conserved in higher plants, including A. thaliana and Oryza sativa. However, it is also assumed that the mechanism differs considerably in detail among species. Here we characterized the clock-controlled CO-FT pathway in Lotus japonicus (a model legume) in comparison with that of A. thaliana. L. japonicus has at least one FT orthologous gene (named LjFTa), which is induced specifically in short-days and complements the mutational lesion of the Arabidopsis FT gene. However, it was speculated that this legume might lack the upstream positive regulator CO. By employing L. japonicus phyB mutant plants, we showed that the photoreceptor mutant displays a phenotype of early flowering due to enhanced expression of LjFTa, suggesting that LjFTa is invovled in the promotion of flowering in L. japonicus. These results are discussed in the context of current knowledge of the flowering in crop legumes such as soybean and garden pea.
    Bioscience Biotechnology and Biochemistry 04/2013; · 1.27 Impact Factor
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    ABSTRACT: Plant circadian clock controls a wide variety of physiological and developmental events, which include the short-days (SDs)-specific promotion of the elongation of hypocotyls during de-etiolation and also the elongation of petioles during vegetative growth. In A. thaliana, the PIF4 gene encoding a phytochrome-interacting basic helix-loop-helix (bHLH) transcription factor plays crucial roles in this photoperiodic control of plant growth. According to the proposed external coincidence model, the PIF4 gene is transcribed precociously at the end of night specifically in SDs, under which conditions the protein product is stably accumulated, while PIF4 is expressed exclusively during the daytime in long days (LDs), under which conditions the protein product is degraded by the light-activated phyB and also the residual proteins are inactivated by the DELLA family of proteins. A number of previous reports provided solid evidence to support this coincidence model mainly at the transcriptional level of the PIF 4 and PIF4-traget genes. Nevertheless, the diurnal oscillation profiles of PIF4 proteins, which were postulated to be dependent on photoperiod and ambient temperature, have not yet been demonstrated. Here we present such crucial evidence on PIF4 protein level to further support the external coincidence model underlying the temperature-adaptive photoperiodic control of plant growth in A. thaliana.
    Plant signaling & behavior 01/2013; 8(3).
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    ABSTRACT: Recent intensive studies of the model plant Arabidopsis thaliana have revealed the molecular mechanisms underlying circadian rhythms in detail. Results of phylogenetic analyses indicated that some of core clock genes are widely conserved throughout the plant kingdom. For another model plant the legume Lotus japonicus, we have reported that it has a set of putative clock genes highly homologous to A. thaliana. Taking advantage of the L. japonicus hairy root transformation system, in this study we characterized the promoter activity of A. thaliana core clock genes CCA1 and PRR5 in heterologous L. japonicus cells and found that the molecular mechanism of circadian rhythm in L. japonicus is compatible with that of A. thaliana.
    Bioscience Biotechnology and Biochemistry 12/2012; · 1.27 Impact Factor
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    ABSTRACT: In Arabidopsis thaliana, the circadian clock regulates the photoperiodic plant growth including the elongation of hypocotyls in a short-days (SDs)-specific manner. The clock-controlled PHYTOCHROME-INTERACTING FACTOR 4 (PIF4) gene encoding a basic helix-loop-helix (bHLH) transcription factor plays crucial roles in this regulation. The SDs-specific elongation of hypocotyls is best explained by accumulation of the active PIF4 proteins at the end of night specifically in SDs due to coincidence between internal (circadian clock) and external (photoperiod) cues. However, this external coincidence model was challenged with the recent finding that the elongation of hypocotyls is markedly promoted at high growth temperature (28 ˚C) even in long-days (LDs), implying that the model to explain the photoperiodic response of plant architecture appears to be conditional on ambient temperature. With regard to this problem, the results of this and previous studies showed that the model holds under a wide range of ambient temperature conditions (16 ˚C to 28 ˚C). We propose that the circadian clock and PIF4-mediated external coincidence mechanism coordinately integrates both of the cues from seasonal changes in photoperiod and temperature to regulate plant growth in natural habitats.
    Plant signaling & behavior 11/2012; 8(2).
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    ABSTRACT: Plant circadian clock generates rhythms with a period close to 24 h, and it controls a wide variety of physiological and developmental events, enabling plants to adapt to ever-changing environmental light conditions. In Arabidopsis thaliana, the clock regulates the diurnal and photoperiodic plant growth including the elongation of hypocotyls and petioles in a time-of-day-specific and short-days (SDs)-specific manner. In this mechanism, the clock-regulated PHYTOCHROME-INTERACTING FACTOR 4 gene encoding a basic helix-loop-helix transcription factor, together with phytochromes (mainly phyB), plays crucial roles. This diurnal and photoperiodic control of plant growth is best explained by the accumulation of the PIF4 protein at the end of nighttime specifically under SDs, due to coincidence between the internal (circadian rhythm) and external (photoperiod) cues. In this model, however, the PIF4-controlled downstream factors are not fully identified, although it has been generally proposed that the auxin-mediated signal transduction is crucially implicated. Here, we identified a set of hormone-associated genes as the specific PIF4-targets implicated in the photoperiodic control of plant growth. They include not only auxin-associated genes (GH3.5, IAA19, IAA29), but also ones associated with other growth-regulating hormones such as brassinosteroids (BR6ox2), gibberellic acids (GAI), ethylene (ACS8), and cytokinin (CKX5). The dawn- and SDs-specific expression profiles of these genes are modified in a set of phyB and clock mutants, both of which compromise the coincidence mechanism. The results of this study suggest that the circadian clock orchestrates a variety of hormone-signaling pathways to regulate the photoperiod-dependent morphogenesis in A. thaliana.
    Plant and Cell Physiology 10/2012; · 4.98 Impact Factor
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    ABSTRACT: In Alabidopsis thaliana, the circadian clock regulates the diurnal and photoperiodic plant growth including the elongation of hypocotyls in a time-of-day-specific and short-days (SDs)-specific manner. The clock-controlled PHYTOCHROME-INTERACTING FACTOR 4 encoding a basic helix-loop-helix (bHLH) transcription factor plays crucial roles in this regulation. PIF4 is transcribed precociously at the end of night in SDs, under which conditions the protein product is stably accumulated, while PIF4 is expressed exclusively during the daytime in long days (LDs), under which conditions the protein product is degraded by the light-activated phyB. The dawn- and SDs-specific elongation of hypocotyls is best explained by the coincident accumulation of the active PIF4 protein during the nighttime before dawn specifically in SDs. However, this coincidence model was challenged with the recent finding that the elongation of hypocotyls is markedly promoted at high growth temperature (i.e., 28 °C) even under LDs in a PIF4-dependent manner. Here, we reconciled the apparently conflicting facts by showing that the transcription of PIF4 occurs precociously at the end of nighttime at 28 °C in LDs, similarly as in SDs. Both the events resulted in the same consequence that a set of PIF4-target genes (ATHB2, GH3.5, IAA19, IAA29, BRox2, GAI, ACS8, and CKX5) was induced accordingly in a time-of-day-specific manner. Taken together, we propose an extended double coincidence mechanism, by which the two environmental cues (i.e., photoperiods and temperatures), both of which vary on a season-to-season basis, are integrated into the same clock and PIF4-mediated output pathway that regulate a hormone-signaling network to fit plant architectures properly to domestic habitats.
    Plant and Cell Physiology 10/2012; · 4.98 Impact Factor
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    ABSTRACT: The circadian clock is an endogenous time-keeping mechanism that enables organisms to adapt to external daily cycles. The clock coordinates biological activities with these cycles, mainly through genome-wide gene expression. However, the exact mechanism underlying regulation of circadian gene expression is poorly understood. Here we demonstrated that an Arabidopsis PSEUDO-RESPONSE REGULATOR 5 (PRR5), which acts in the clock genetic circuit, directly regulates expression timing of key transcription factors involved in clock-output pathways. A transient expression assay and ChIP-quantitative PCR assay using mutated PRR5 indicated that PRR5 associates with target DNA through binding at the CCT motif in vivo. ChIP followed by deep sequencing coupled with genome-wide expression profiling revealed the direct-target genes of PRR5. PRR5 direct-targets include genes encoding transcription factors involved in flowering-time regulation, hypocotyl elongation, and cold-stress responses. PRR5-target gene expression followed a circadian rhythm pattern with low, basal expression from noon until midnight, when PRR9, PRR7, and PRR5 were expressed. ChIP-quantitative PCR assays indicated that PRR7 and PRR9 bind to the direct-targets of PRR5. Genome-wide expression profiling using a prr9 prr7 prr5 triple mutant suggests that PRR5, PRR7, and PRR9 repress these targets. Taken together, our results illustrate a genetic network in which PRR5, PRR7, and PRR9 directly regulate expression timing of key transcription factors to coordinate physiological processes with daily cycles.
    Proceedings of the National Academy of Sciences 10/2012; 109(42):17123-8. · 9.81 Impact Factor
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    ABSTRACT: Sessile plants must continuously adjust their growth and development to optimize photosynthetic activity under ever-fluctuating light conditions. Among such light responses in plants, one of the best-characterized events is the so-called shade avoidance, for which a low ratio of the red (R):far-red (FR) light intensities is the most prominent stimulus. Such shade avoidance responses enable plants to overtop their neighbors, thereby enhancing fitness and competitiveness in their natural habitat. Considerable progress has been achieved during the last decade in understanding the molecular mechanisms underlying the shade avoidance responses in the model rosette plant, Arabidopsis thaliana. We characterize here the fundamental aspects of the shade avoidance responses in the model legume, Lotus japonicus, based on the fact that its phyllotaxis (or morphological architecture) is quite different from that of A. thaliana. It was found that L. japonicus displays the characteristic shade avoidance syndrome (SAS) under defined laboratory conditions (a low R:FR ratio, low light intensity, and low blue light intensity) that mimic the natural canopy. In particular, the outgrowth of axillary buds (i.e., both aerial and cotyledonary shoot branching) was severely inhibited in L. japonicus grown in the shade. These results are discussed with special emphasis on the unique aspects of SAS observed with this legume.
    Bioscience Biotechnology and Biochemistry 11/2011; 75(11):2148-54. · 1.27 Impact Factor
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    ABSTRACT: The developmental programs of Physcomitrella patens, a basal lineage of land plants, are regulated by phytohormones and light-signaling responses. In this study, our attention was focused on the HY5-family of transcription factors, which are known to play important roles immediately downstream of photoreceptors during the early photomorphogenesis of Arabidopsis thaliana. We retrieved two HY5-homologs, named PpHY5a and PpHY5b, from the whole genome sequence database of P. patens. Arabidopsis transgenic plants overproducing the basic leucine zipper (bZIP) domain of PpHY5a exhibited a phenotype of short hypocotyls, suggesting a functional relationship between PpHY5 and Arabidopsis HY5. A loss-of-function Δhy5a Δhy5b double mutant was defective in the vigorous protrusion of caulonema cells from the protonema networks of P. patens under light and dark conditions. These results suggest that the function of HY5-homologs in P. patens is evolutionarily conserved, and is implicated in a process of caulonema development.
    Bioscience Biotechnology and Biochemistry 08/2011; 75(8):1533-9. · 1.27 Impact Factor
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    ABSTRACT: The plant circadian clock generates rhythms with a period close to 24 h, and it controls a wide variety of physiological and developmental events. Among clock-controlled developmental events, the best characterized is the photoperiodic control of flowering time, which is mediated through the CONSTANS (CO)-FLOWERING LOCUS T (FT) pathway in Arabidopsis thaliana. The clock also regulates the diurnal plant growth including the elongation of hypocotyls in a short day (SDs)-specific manner. In this mechanism, phytochromes (mainly phyB) and the PHYTOCHROME-INTERACTING FACTOR4 (PIF4) and PIF5, encoding phytochrome-interacting basic helix-loop-helix (bHLH) transcription factors, play crucial roles. The time of day-specific and photoperiodic control of hypocotyl elongation is best explained by the accumulation of the PIF4 and PIF5 proteins during night-time before dawn, especially under SDs, due to coincidence between the internal (circadian rhythm) and external (photoperiod) time cues. However, the PIF4- and/or PIF5-controlled downstream factors have not yet been identified. Here, we provide evidence that ARABIDOPSIS THALIANA HOMEOBOX PROTEIN2 (ATHB2), together with auxin-inducible IAA29, is diurnally expressed with a peak at dawn under the control of PIF4 and PIF5 specifically in SDs. This coincidentally expressed transcription factor serves as a positive regulator for the elongation of hypocotyls. The expression profiles of ATHB2 were markedly altered in certain clock and phytochrome mutants, all of which show anomalous phenotypes with regard to the photoperiodic control of hypocotyl elongation. Taken together, we propose that an external coincidence model involving the clock-controlled PIF4/PIF5-ATHB2 pathway is crucial for the diurnal and photoperiodic control of plant growth in A. thaliana.
    Plant and Cell Physiology 06/2011; 52(8):1315-29. · 4.98 Impact Factor
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    Daisuke Hagiwara, Takeshi Mizuno, Keietsu Abe
    Current Genetics 06/2011; 57(3):223-4. · 2.41 Impact Factor
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    ABSTRACT: Physcomitrella patens has four homologs of the pseudo-response regulator involved in the circadian clock mechanism in seed plants. To gain insight into their function, Arabidopsis transgenic lines misexpressing PpPRR2 were constructed. Phenotypic analysis of the transformants with reference to clock-related gene expression and photoperiodic responses revealed that heterologous expression of the moss PpPRR2 gene modifies the intrinsic mechanism underlying the circadian clock in Arabidopsis, suggesting that PpPRR2 serves as a clock component in P. patens.
    Bioscience Biotechnology and Biochemistry 05/2011; 75(4):786-9. · 1.27 Impact Factor
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    ABSTRACT: The pseudo-response regulators (PRRs) are the circadian clock component proteins in the model dicot Arabidopsis thaliana. They contain a receiver-like domain (RLD) similar to the receiver domains of the RRs in the His-Asp phosphorelay system, but the RLDs lack the phosphoacceptor aspartic acid residue invariably conserved in the receiver domains. To study the evolution of PRR genes in plants, here we characterize their homologue genes, PpPRR1, PpPRR2, PpPRR3 and PpPRR4, from the moss Physcomitrella patens. In the phylogenetic analysis, PpPRRs cluster together, sister to an angiosperm PRR gene subfamily, illustrating their close relationships with the angiosperm PRRs. However, distinct from the angiosperm sequences, the RLDs of PpPRR2/3/4 exhibit a potential phosphoacceptor aspartic acid-aspartic acid-lysine (DDK) motif. Consistently, the PpPRR2 RLD had phosphotransfer ability in vitro, suggesting that PpPRR2 functions as an RR. The PpPRR1 RLD, on the other hand, shows a partially diverged DDK motif, and it did not show phosphotransfer ability. All PpPRRs were expressed in a circadian and light-dependent manner, with differential regulation between PpPRR2/4 and PpPRR1/3. Altogether, our results illustrate that PRRs originated from an RR(s) and that there are intraspecific divergences among PpPRRs. Finally, we offer scenarios for the evolution of the PRR family in land plants.
    DNA Research 01/2011; 18(1):39-52. · 4.43 Impact Factor
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    ABSTRACT: In the model seed plant Arabidopsis thaliana, a sub-family of B-box containing transcriptional factors (BBXs), which is classified in the BBX-IV group based on the domain structure, contains two tandem B-box domains and plays crucial roles in early photomorphogenesis under the control of blue light receptors, cry1 and cry2. The results of an examination of light responsiveness of representative Physcomitrella BBX-IV genes and their heterologous expression in Arabidopsis suggested that the light signaling-related characteristics of the BBX-IV subfamily are evolutionarily conserved in a moss, which is a basal lineage of land plants.
    Bioscience Biotechnology and Biochemistry 01/2011; 75(10):2037-41. · 1.27 Impact Factor
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    Daisuke Hagiwara, Takeshi Mizuno, Keietsu Abe
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    ABSTRACT: Ssk1- and Skn7-type response regulators are widely conserved in fungal His-Asp phosphorelay (two-component) signaling systems. SrrA, a Skn7-type RR of Aspergillus nidulans, is implicated not only in oxidative stress responses but also in osmotic adaptation, conidia production (asexual development), inhibition by fungicides, and cell wall stress resistance. Here, we characterized SrrA, focusing on the role of the conserved aspartate residue in the receiver domain, which is essential for phosphorelay function. We constructed strains carrying an SrrA protein in which aspartate residue D385 was replaced with either asparagine (N) or alanine (A). These mutants exhibited normal conidiation and partial oxidative stress resistance. In osmotic adaptation, mutants with substitution at SrrA D385 showed as much sensitivity as ΔsrrA strains, suggesting that SrrA plays a role in osmotic stress adaptation in a phosphorelay-dependent manner. The SrrA D385 substitution mutants showed significant resistance to fungicides and cell wall stresses. These results together led us to conclude that the conserved aspartate residue has a substantial impact on SrrA function, and that SrrA plays a role in several aspects of cellular function via His-Asp phosphorelay circuitry in Aspergillus nidulans.
    Current Genetics 01/2011; 57(2):103-14. · 2.41 Impact Factor

Publication Stats

10k Citations
1,011.19 Total Impact Points

Institutions

  • 1985–2014
    • Nagoya University
      • • Graduate School of Bio-Agricultural Sciences
      • • Department of Biological Mechanisms and Functions
      Nagoya, Aichi, Japan
  • 2011
    • Chuo University
      • Department of Biological Sciences
      Tokyo, Tokyo-to, Japan
  • 2007–2011
    • Tohoku University
      • Graduate School of Agricultural Science
      Sendai-shi, Miyagi-ken, Japan
    • Osaka Prefecture University
      • Graduate School of Life and Environmental Sciences
      Sakai, Osaka-fu, Japan
  • 1996–2009
    • Nara Institute of Science and Technology
      • • Graduate School of Biological Sciences
      • • Research and Education Center for Genetic Information
      Ikuma, Nara, Japan
  • 2004
    • Sony Corporation
      Edo, Tōkyō, Japan
    • Lawrence Berkeley National Laboratory
      Berkeley, California, United States
  • 2001–2004
    • Waseda University
      • Department of Applied Physics
      Tokyo, Tokyo-to, Japan